Effect of Long-term Fasting on the Use of Fatty Acids as Trophic Markers in the Opossum Shrimp Mysis relicta—A Laboratory Study

Fatty acids are potential trophic markers to trace feeding relationships in aquatic ecosystems primarily because lipid reserves of organisms broadly reflect dietary sources of lipids and can therefore provide information on the availability of key fatty acids in the food web. However, the use of fatty acids for such applications may be constrained by the degree to which the fatty acid composition of organisms is obscured by factors other than straightforward uptake from the diet. Thus, we studied the effect of long-term fasting, under controlled laboratory conditions, on the lipid content and fatty acid composition of field-caught Mysis relicta. Periods of 3 to 6 weeks are required to induce clear effects of fasting in M. relicta. Relative proportions of docosahexaenoic acid (DHA; 22:6n-3) and arachidonic acid (ARA; 20:4n-6) increased with decreasing lipid contents during fasting. DHA proportions in the total fatty acid composition of fasting M. relicta were significantly higher in comparison to field-caught animals. M. relicta with high proportions of DHA (> 25%) and with low lipid contents (< 14% of dry weight) can be clearly identified as fasting animals. Such thresholds will help to improve the validity of fatty acids in trophic studies.     

Spatial variation in RNA:DNA ratios of Diporeia spp. in the Great Lakes region

Over the past two decades, Diporeia in all of the Laurentian Great Lakes, except Superior, have declined dramatically. These declines have seemingly coincided with expansion of invasive Dreissena polymorpha and D. bugensis, however the exact mechanisms underlying decreasing Diporeia densities are obscure. We explored the use of RNA:DNA (R/D) ratios as a conditional index for Diporeia by experimentally demonstrating that Diporeia R/D responds to periods of starvation. Moreover, during 2008–2009 we collected Diporeia from throughout the Great Lakes and Cayuga Lake (New York, USA), and used R/D ratios to index condition of these in situ collected animals. We evaluated spatial and temporal variation of nucleic acid indices using classification and regression tree (CART) analysis with a suite of environmental variables included as potential predictors. Diporeia R/D of in situ collected specimens exhibited pronounced spatial and temporal variation, but multiple CART models described only a small amount of this variation. While we observed some variation in Diporeia R/D among lakes, nucleic acid ratios appeared to respond weakly to Diporeia population characteristics and local environmental conditions. Specifically, CART analyses revealed that Diporeia R/D was particularly low at extreme depths, and interestingly, Diporeia nucleic acids were not strongly associated with the presence of dreissenids. In summary, while a limited amount of variation in Diporeia R/D was attributable to environmental conditions, the majority of Diporeia R/D variation was unaccounted for. Hence, the causative factors underlying spatio-temporal variation of Diporeia R/D and the mechanistic reasons for Diporeia declines in the Great Lakes remain largely unknown.     

Diet effects on lipid composition,somatic growth potential,and survival of the benthic amphipod Diporeia spp.

The benthic amphipod Diporeia represents a crucial trophic link that conveys vital nutrients and energy to predators at higher trophic levels. The current decline of Diporeia populations, mostly in the North American Great Lakes, may, in part, be related to concurrent declines in food quantity and/or quality. We hypothesized that somatic growth and survival of Diporeia would be positively related to dietary supply and subsequent retention of polyunsaturated fatty acids (PUFA); a class of chemicals known to affect diet quality. We examined how different algal PUFA concentrations in; a) Ankistrodesmus falcatus (Chlorophyta), b) a naturally occurring diatom assemblage from Lake Ontario, c) a non-toxic strain of Microcystis aeruginosa (Cyanophyta), and, d) fasting for 30 d, affected PUFA concentrations, somatic growth, and survival of Diporeia. Total PUFA concentrations were significantly higher in A. falcatus than in diatoms and Microcystis, but only diatoms contained considerable amounts of eicosapentaenoic (EPA) and docosahexaenoic acid (DHA). EPA, DHA, and arachidonic acid (ARA) were highly retained in Diporeia even in the absence of dietary supply with ARA being the most efficiently bioaccumulated PUFA. Survivorship of Diporeia ranged from 60% (diatom-fed), 68% (A. falcatus-fed), to 70% (fasting treatment), but was 0% in the M. aeruginosa diet treatment. Nucleic acid ratios (RNA:DNA), commonly used as proxies for somatic growth potential, were highest in Diporeia feeding on diatoms and lowest in fasting animals. We conclude that overall condition of Diporeia improved with dietary access to EPA and DHA, but survival was not related to this food quantity and/or quality.     

Gastric evacuation rate,index of fullness,and daily ration of Lake Michigan slimy (Cottus cognatus) and deepwater sculpin (Myoxocephalus thompsonii)

Accurate estimates of fish consumption are required to understand trophic interactions and facilitate ecosystem-based fishery management. Despite their importance within the food-web, no method currently exists to estimate daily consumption for Great Lakes slimy (Cottus cognatus) and deepwater sculpin (Myoxocephalus thompsonii). We conducted experiments to estimate gastric evacuation (GEVAC) and collected field data from Lake Michigan to estimate index of fullness [(g prey/g fish weight)100%) to determine daily ration for water temperatures ranging 2–5 °C, coinciding with the winter and early spring season. Exponential GEVAC rates equaled 0.0115/h for slimy sculpin and 0.0147/h for deepwater sculpin, and did not vary between 2.7 °C and 5.1 °C for either species or between prey types (Mysis relicta and fish eggs) for slimy sculpin. Index of fullness varied with fish size, and averaged 1.93% and 1.85% for slimy and deepwater sculpins, respectively. Maximum index of fullness was generally higher (except for the smallest sizes) for both species in 2009–2010 than in 1976 despite reductions in a primary prey, Diporeia spp. Predictive daily ration equations were derived as a function of fish dry weight. Estimates of daily consumption ranged from 0.2 to 0.8% of their body weight, which was within the low range of estimates from other species at comparably low water temperatures. These results provide a tool to estimate the consumptive demand of sculpins which will improve our understanding of benthic offshore food webs and aid in management and restoration of these native species in the Great Lakes.     

A short-term look at potential changes in Lake Michigan slimy sculpin diets

Diporeia hoyi and Mysis relicta are the most important prey items of slimy sculpins (Cottus cognatus) in the Great Lakes. Slimy sculpins were collected from dreissenid-infested bottoms off seven Lake Michigan ports at depths of 27–73 m in fall 2003 to study their lake-wide diets. Relatively large dreissenid biomass occurred at depths of 37- and 46-m. Quagga mussels (Dreissena bugnesis) composed at least 50% of dreissenid biomass at Manistique, Saugatuck, and Sturgeon Bay. Mysis accounted for 82% of the sculpin diet by dry weight at eastern Lake Michigan while Diporeia composed 54–69% of the diet at western Lake Michigan and dominated the diets of slimy sculpins at all sites deeper than 46 m. In northern Lake Michigan, this diet study in new sites showed that slimy sculpin consumed more prey with low energy contents, especially chironomids, than Mysis and Diporeia in shallow sites (depth <55 m). We recommend diet studies on sedentary benthic fishes to be conducted along perimeters of the Great Lakes to observe changes in their diets that may be impacted by changing benthic macroinvertebrate communities.     

Predation by Mysis Relicta on Pontoporeia Hoyi: A Food Chain Link of Potential Importance in the Great Lakes

Sediment cores composed of benthic microorganisms and the natural assemblage of benthic fauna from Lakes Michigan and Erie were maintained intact at 4° C in a laboratory incubator for 10 weeks. Visual inspection of the microcosms showed that the opossum shrimp Mysis relicta, an omnivorous planktivore, frequently preyed on the burrowing amphipod Pontoporeia hoyi, a benthic detritivore; a previously unreported predator/prey relationship. Predation by Mysis on Pontoporeia demonstrates that Mysis can utilize not only the plankton carbon pool but the benthic/detrital carbon pool as well. This relationship may represent an important link in the food web of the Great Lakes, since Pontoporeia is the dominant macrobenthic organism in the lakes and both organisms are an important food source for immature salmonids and other fish species in the Great Lakes.     

What's in a name? Taxonomy and nomenclature of invasive gobies in the Great Lakes and beyond

The species identities, scientific names, and relationships of Eurasian gobies that invaded the Laurentian Great Lakes – and other species that are predicted to invade in the future – are evaluated here using recently resolved DNA characters. The Round Goby and the Freshwater Tubenose Goby entered the Great Lakes ca. 1990 via ballast water originating from Black Sea ports. The Round Goby spread extensively throughout the Great Lakes and adjacent rivers, whereas the Freshwater Tubenose Goby recently began to expand its range. Both species also are widely invasive in Eurasia, dispersing via canals and shipping. Several of their relatives – the Monkey, Racer, and Bighead gobies – also are invasive in Eurasia, and are predicted to invade the Great Lakes. We discuss results from phylogenetic analyses of DNA sequences from 4 mitochondrial and nuclear gene regions, and provide a revision of their scientific nomenclature. The Freshwater Tubenose Goby was redefined as Proterorhinus semilunaris, which is markedly different and distinctive from the Marine Tubenose Goby Proterorhinus marmoratus. The genus Neogobius, as formerly defined, contained multiple evolutionary lineages and incorrect scientific names. We thus restricted Neogobius to just 4 species—including the Round Goby Neogobius melanostomus and the Black Sea Monkey Goby Neogobius fluviatilis. Several previously recognized subgenera, which were incorrectly grouped in Neogobius, were elevated to the level of genera. Notably, the Racer Goby became Babka gymnocephalus and the Bighead Goby now is Ponticola kessleri. These changes made the names consistent with their true relationships and species characters, which are essential for identifying and characterizing these gobies in invasive and native habitats.     

The establishment of the nuisance cyanobacteria Lyngbya wollei in Lake St. Clair and its potential to harbor fecal indicator bacteria

Lyngbya wollei is a filamentous cyanobacterium which forms large nuisance mats and has infested eastern and southeastern U.S. Lakes and reservoirs for over 100 years. Lyngbya was recently identified in the Great Lakes system in the St. Lawrence River, and Western Lake Erie. Here we report on large deposits of L. wollei washing onshore at a popular recreational beach in Lake Saint Clair, part of the Great Lakes system. The amount of L. wollei deposited on shore was quantified and evaluated for the presence of fecal indicator bacteria (FIB). High concentrations of Escherichia coli, enterococci and Clostridium perfringens were found in the L. wollei in nearshore waters. The densities of E. coli (MPN), enterococci (MPN) and C. perfringens (CFU) attached to L. wollei averaged 3.5, 3.2 and 3.2 log/g, respectively. In contrast, nearshore waters contained nearly 10 times less FIB, averaging 2.6, 2.4 and 2.6 log/100 ml of E. coli (MPN), enterococci (MPN) and C. perfringens (CFU), respectively. DNA fingerprint analysis was used to examine the population structure of E. coli isolates obtained from L. wollei mats. The L. wollei-borne E. coli strains were genetically diverse, suggesting a causal relationship between E. coli and L. wollei. Results from this study indicate that in addition to the macroalga such as Cladophora, cyanobacteria like L. wollei also harbor FIB, potentially impacting water quality and human health in the Great Lakes.     

Changes in Fecundity and Egg Lipid Content of Lake Whitefish (Coregonus clupeaformis) in the Upper Laurentian Great Lakes between 1986–87 and 2003–05

Lake whitefish (Coregonus clupeaformis Mitchill), an important commercial species in the Laurentian Great Lakes, have experienced decreased growth and condition in regions of the upper Great Lakes over the past 20 years. Increases in lake whitefish density and decreases in the density of Diporeia spp., an energy rich and historically important part of the lake whitefish diet, have been implicated in the recent declines in lake whitefish growth and condition. The goal of this study was to describe lake whitefish fecundity, egg lipid content, and total ovary lipid content in selected regions of Lakes Huron, Michigan, and Superior in 1986–87 and 2003–05, two time periods with different lake whitefish and Diporeia densities. Under conditions of high lake whitefish density and low Diporeia density, female lake whitefish in the upper Laurentian Great Lakes generally produced fewer eggs. Egg lipid content was higher in 2003–05 than in 1986–87 at all sites, regardless of changes in lake whitefish or Diporeia densities. Total ovary lipid content and lake whitefish abundance were inversely related, while there was no significant relationship between total ovary lipid content and Diporeia density. The amount of energy that lake whitefish invested in egg production was more closely associated with lake whitefish abundance than with Diporeia density. This study provides evidence that recent changes in production dynamics of Great Lakes lake whitefish have not been driven solely by declines in Diporeia but have been significantly influenced by lake whitefish abundance.     

Metazoan Parasites of Introduced Round and Tubenose Gobies in the Great Lakes: Support for the “Enemy Release Hypothesis”

Recent invasion theory has hypothesized that newly established exotic species may initially be free of their native parasites, augmenting their population success. Others have hypothesized that invaders may introduce exotic parasites to native species and/or may become hosts to native parasites in their new habitats. Our study analyzed the parasites of two exotic Eurasian gobies that were detected in the Great Lakes in 1990: the round goby Apollonia melanostoma and the tubenose goby Proterorhinus semilunaris. We compared our results from the central region of their introduced ranges in Lakes Huron, St. Clair, and Erie with other studies in the Great Lakes over the past decade, as well as Eurasian native and nonindigenous habitats. Results showed that goby-specific metazoan parasites were absent in the Great Lakes, and all but one species were represented only as larvae, suggesting that adult parasites presently are poorly-adapted to the new gobies as hosts. Seven parasitic species are known to infest the tubenose goby in the Great Lakes, including our new finding of the acanthocephalan Southwellina hispida, and all are rare. We provide the first findings of four parasite species in the round goby and clarified two others, totaling 22 in the Great Lakes—with most being rare. In contrast, 72 round goby parasites occur in the Black Sea region. Trematodes are the most common parasitic group of the round goby in the Great Lakes, as in their native Black Sea range and Baltic Sea introduction. Holarctic trematode Diplostomum spathaceum larvae, which are one of two widely distributed species shared with Eurasia, were found in round goby eyes from all Great Lakes localities except Lake Huron proper. Our study and others reveal no overall increases in parasitism of the invasive gobies over the past decade after their establishment in the Great Lakes. In conclusion, the parasite “load” on the invasive gobies appears relatively low in comparison with their native habitats, lending support to the “enemy release hypothesis.”     

Laboratory-derived temperature preference and effect on the feeding rate and survival of juvenile Hemimysis anomala

Hemimysis anomala is a warm-water mysid that invaded the Great Lakes region in 2006 and has since rapidly spread throughout the basin. We conducted three laboratory experiments to better define the temperature preference, tolerance limits, and temperature effects on feeding rates of juvenile Hemimysis, using individuals acclimated to mid (16 °C) and upper (22 °C) preferred temperature values previously reported for the species. For temperature preference, we fit a two-parameter Gaussian (μ, σ) function to the experimental data, and found that the peak values (μ, interpreted as the preference temperature) were 22.0 °C (SE 0.25) when acclimated to 16 and 21.9 °C (SE 0.38) when acclimated to 22 °C, with the σ-values of the curves at 2.6 and 2.5 °C, respectively. No mysids were observed in temperatures below 10 or above 28 °C in these preference experiments. In short-term tolerance experiments for temperatures between 4 and 32 °C, all mysids died within 8 h at 30.2 °C for 16 °C acclimated mysids, and at 31.8 °C for 22 °C acclimated mysids. No lower lethal limit was found. Feeding rates increased with temperature from an average of 4 Bosmina eaten per hour at 5 °C to 19 Bosmina eaten per hour at 27 °C. The results of our experiments indicate an optimal temperature for Hemimysis between 21 and 27 °C, which corresponds with temperatures during periods of high population growth in the field. These results contribute a better understanding of this species' biological response to temperature that will help guide field studies and inform bioenergetics modeling.     

Alewife Mortality,Condition, and Immune Response to Prolonged Cold Temperatures

Alewife, Alosa pseudoharengus, have been recognized for several decades as one of the most important forage fish in the Laurentian Great Lakes. Although massive alewife die-offs have regularly been observed throughout the Great Lakes and other inland lakes, little substantive information is available regarding physiological mechanisms associated with adult alewife mortality. Long-term field surveys have shown a correlation between cold winter temperatures, poor condition and adult alewife mortality. In this study, adult alewife were raised in replicate pond systems and subjected to contrasting cold temperatures (4 and < 2°C) representing mild and severe winter conditions. We evaluated alewife mortality, condition and immune response to these temperatures. In contrast to our expectations, alewife exposed to mild and severe winter temperatures showed no difference in mortality or condition (measured as the ratio of dry to wet weight). Survival of alewife held in ponds with mild winter conditions (∼ 4°C) was similar to that of alewife exposed to prolonged periods (more than six weeks) of temperatures < 2°C. This result contrasts with previous observations indicating that alewife cannot tolerate temperatures < 3°C. Circulating lymphocytes from alewife exposed to severe winter temperatures were significantly lower in number (∼40%) compared to fish experiencing milder° winter conditions, suggesting sub-lethal immunosuppression in response to the colder winter temperatures. Although colder winter temperatures did not directly induce alewife mortality, these results suggest that winter conditions that result in colder water temperatures can produce immunosuppression, thereby increasing alewife susceptibility to disease and mortality.     

Early observations on an emerging Great Lakes invader Hemimysis anomala in Lake Ontario

Hemimysis anomala, a Ponto-Caspian littoral mysid, is an emerging Great Lakes invader that was discovered in Lakes Michigan and Ontario in 2006. Similar to the native mysid Mysis diluviana, Hemimysis exhibits a diel vertical migration pattern but generally inhabits shallower and warmer waters than M. diluviana. Because basic information on the distribution, habitat use, and biology of Hemimysis in the Great Lakes is scarce, the potential for food web disruption by Hemimysis cannot easily be predicted. Preliminary observations indicate widespread invasion of Hemimysis in Lake Ontario. In this study, we confirm the presence of Hemimysis at sites spanning the northern and southern shores of Lake Ontario and the presence of the individuals during winter months. In one horizontal tow in November 2007, over 26,000 individuals were collected with a length range of 4.4 to 9.0 mm and an average caloric density of 611 cal/g wet weight. The most effective methods for sampling Hemimysis were horizontal tows with either a zooplankton net in the water column or a benthic sled near the lake bottom. Although more quantitative data on the life history and distribution of this species is necessary, our preliminary observations support the prediction that the potential for Hemimysis to impact the nearshore food web in Lake Ontario appears high.     

Distribution,seasonality and putative origin of the non-native red alga Bangia atropurpurea (Bangiales,Rhodophyta) in the Laurentian Great Lakes

Bangia atropurpurea was first observed in Lake Erie in 1964 and subsequently spread to the lower Laurentian Great Lakes by the mid to late 1970s. The present study was initiated to examine the recent distribution of B. atropurpurea in the Great Lakes, the seasonal variation of the alga and the putative origin of this species based on DNA sequence analysis. From surveys in 1995 and 2002, this species has clearly spread, with newly identified populations observed in Lakes Huron, Michigan, Georgian Bay and the St. Lawrence River. Morphological analyses showed that Great Lakes populations from individual lakes or neighboring populations did not group together in cluster analyses. Correlation analysis, however, revealed significant relationships between the presence or absence of Bangia among the studied sites with pH and specific conductance as those locations that had stable populations had a mean pH and conductance of 8.2 and 353 μS·cm^− 1 respectively. Seasonal variation in morphology of a population from Burlington, Ontario (Lake Ontario, Canada) was examined monthly for one year. Maximum filament length occurred in April, with the greatest diameter and archaeospore production observed in May. Significant correlations were also noted for many morphological characteristics with water temperature, population height on the shoreline relative to the waterline and total phosphorus. Collections of B. atropurpurea analyzed from the Great Lakes were observed to be identical in sequence to collections of European freshwater Bangia in the cox2–3 gene spacer, the nuclear internal transcribed spacers (ITS 1 and 2) and the 5.8S rRNA gene (between the small and large subunits of the rRNA gene). These results suggest a recent European origin; however, further global collections of B. atropurpurea and microsatellite analyses are necessary to confirm this hypothesis.     

Associations between Breeding Marsh Bird Abundances and Great Lakes Hydrology

We used Great Lakes hydrologic data and bird monitoring data from the Great Lakes Marsh Monitoring Program from 1995–2002 to: 1) evaluate trends and patterns of annual change in May-July water levels for Lakes Ontario, Erie, and Huron-Michigan, 2) report on trends of relative abundance for birds breeding in Great Lakes coastal marshes, and 3) correlate basin-wide and lake-specific annual indices of bird abundance with Great Lakes water levels. From 1995–2002, average May, June, and July water levels in all lake basins showed some annual variation, but Lakes Erie and Huron-Michigan had identical annual fluctuation patterns and general water level declines. No trend was observed in Lake Ontario water levels over this period. Abundance for five of seven marsh birds in Lake Ontario wetlands showed no temporal trends, whereas abundance of black tern (Chlidonias niger) declined and that of swamp sparrow (Melospiza georgiana) increased from 1995–2002. In contrast, abundances of American coot (Fulica americana), black tern, common moorhen (Gallinula chloropus), least bittern (Ixobrychus exilis), marsh wren (Cistorthorus palustris), pied-billed grebe (Podilymbus podiceps), sora (Porzana carolina), swamp sparrow, and Virginia rail (Rallus limicola) declined within marshes at Lakes Erie and Huron/Michigan from 1995–2002. Annual abundances of several birds we examined showed positive correlations with annual lake level changes in non-regulated Lakes Erie and Huron/Michigan, whereas most birds we examined in Lake Ontario coastal wetlands were not correlated with suppressed water level changes of this lake. Overall, our results suggest that long-term changes and annual water level fluctuations are important abiotic factors affecting abundance of some marsh-dependent birds in Great Lakes coastal marshes. For this reason, wetland bird population monitoring initiatives should consider using methods in sampling protocols, or during data analyses, to account for temporal and spatial components of hydrologic variability that affect wetlands and their avifauna.     

Genetic Analysis of the Chinese Mitten Crab (Eriocheir sinensis) Introduced to the North American Great Lakes and St. Lawrence Seaway

The Chinese mitten crab (Eriocheir sinensis) is an invasive organism of concern, with established non-native populations in Europe and California, USA. The species is thought to pose a risk to other North American waterways, including the Great Lakes and St. Lawrence Seaway. Since 1965, there have been sixteen confirmed adult E. sinensis caught in the North American Great Lakes or adjoining waterways. Analysis of their mitochondrial DNA sequence variation for part of the cytochrome c oxidase subunit I gene discerned three haplotypes among seven individuals (caught between 1973 and 2005), identical to common haplotypes in Europe. Analysis of mitochondrial haplotype frequencies and shipping patterns suggests that E. sinensis has been introduced to the Great Lakes from Europe, although we are unable to preclude native Asian populations as putative sources. The species is catadromous, migrating between salt and fresh water to complete its life cycle. This trait makes it unlikely that E. sinensis will establish a breeding population in the Great Lakes proper, which are separated from saltwater by a considerable distance and significant instream barriers such as waterfalls and navigation locks. However, the recent discovery of two confirmed mitten crabs in the St. Lawrence River, which could be more readily colonized, underscores the risk posed by the repeated introduction of this species into the Great Lakes and St. Lawrence Seaway.     

Changes in mid-summer water temperature and clarity across the Great Lakes between 1968 and 2002

In recent decades, three important events have likely played a role in changing the water temperature and clarity of the Laurentian Great Lakes: 1) warmer climate, 2) reduced phosphorus loading, and 3) invasion by European Dreissenid mussels. This paper compiled environmental data from government agencies monitoring the middle and lower portions of the Great Lakes basin (lakes Huron, Erie and Ontario) to document changes in aquatic environments between 1968 and 2002. Over this 34-year period, mean annual air temperature increased at an average rate of 0.037 °C/y, resulting in a 1.3 °C increase. Surface water temperature during August has been rising at annual rates of 0.084 °C (Lake Huron) and 0.048 °C (Lake Ontario) resulting in increases of 2.9 °C and 1.6 °C, respectively. In Lake Erie, the trend was also positive, but it was smaller and not significant. Water clarity, measured here by August Secchi depth, increased in all lakes. Secchi depth increased 1.7 m in Lake Huron, 3.1 m in Lake Ontario and 2.4 m in Lake Erie. Prior to the invasion of Dreissenid mussels, increases in Secchi depth were significant (p < 0.05) in lakes Erie and Ontario, suggesting that phosphorus abatement aided water clarity. After Dreissenid mussel invasion, significant increases in Secchi depth were detected in lakes Ontario and Huron.     

Genetic and Biogeographic Relationships of the Invasive Round (Neogobius melanostomus) and Tubenose (Proterorhinus marmoratus) Gobies in the Great Lakes Versus Eurasian Populations

Population genetic structure and systematic relationships were investigated for two exotic fishes introduced to the Great Lakes in 1990, the round goby Neogobius melanostomus and the tubenose goby Proterorhinus marmoratus, using DNA sequences from the left domain of the mitochondrial DNA control region. Samples of round gobies were compared from different sites in the Great Lakes, an introduced population from the Gulf of Gdansk in Poland, and a native population from the northern Black Sea. The round goby was characterized by relatively high genetic variability, and 17 haplotypes were identified from 64 individuals. Levels of genetic variation for the round goby were similar in the invasive and native sampling sites, suggesting relatively large founding populations and lack of bottlenecks. The northern Black Sea was eliminated as a probable founding source for both the Great Lakes and the exotic population in Poland. Substitutions in the left domain of the control region revealed significant differences among samples from the Great Lakes and Eurasia, and between Lakes Erie and St. Clair, suggesting non-random mating. No variation was detected in the tubenose goby population in the Great Lakes, which has been less successful in terms of spread and population growth. A molecular clock calibration suggested that the genera Neogobius and Proterorhinus diverged about 5.2 ± 1.0 million years ago, apparently separating from a common ancestor shared with Gobius during the isolation of the Paratethys basin from the Mediterranean Tethys Sea.     

Daphnia lumholtzi Sars a new alien species in Lake St. Clair

Daphnia lumholtzi Sars, an exotic tropical/subtropical cladoceran from Australia, southeast Asia, and Africa, was newly found in Lake St. Clair in a vertical tow sample taken at 3 m depth on 25 July 2007. The species was previously found in 1990/1991 in some reservoirs in the southern United States from where it colonized many waters north to the Great Lakes. In 1999, it was found in Lake Erie. This cladoceran had a density of 117 individuals/m³ when we collected it from Lake St. Clair and it was represented by both females (95.12 %) and males (4.88 %). It seems that D. lumholtzi will continue to expand its distribution area in the Great Lakes.     

Historical changes and current status of crayfish diversity and distribution in the Laurentian Great Lakes

Despite increasing recognition of the importance of invertebrates, and specifically crayfish, to nearshore food webs in the Laurentian Great Lakes, past and present ecological studies in the Great Lakes have predominantly focused on fishes. Using data from many sources, we provide a summary of crayfish diversity and distribution throughout the Great Lakes from 1882 to 2008 for 1456 locations where crayfish have been surveyed. Sampling effort was greatest in Lake Michigan, followed by lakes Huron, Erie, Superior, and Ontario. A total of 13 crayfish species occur in the lakes, with Lake Erie having the greatest diversity (n = 11) and Lake Superior having the least (n = 5). Five crayfish species are non-native to one or more lakes. Because Orconectes rusticus was the most widely distributed non-native species and is associated with known negative impacts, we assessed its spread throughout the Great Lakes. Although O. rusticus has been found for over 100 years in Lake Erie, its spread there has been relatively slow compared to that in lakes Michigan and Huron, where it has spread most rapidly since the 1990s and 2000, respectively. O. rusticus has been found in both lakes Superior and Ontario for 22 and 37 years, respectively, and has expanded little in either lake. Our broad spatial and temporal assessment of crayfish diversity and distribution provides a baseline for future nearshore ecological studies, and for future management efforts to restore native crayfish and limit non-native introductions and their impact on food web interactions.